Floristic relationships among inland swamp forests of Southeastern and Central-Western Brazil

Relações florísticas entre florestas paludosas interioranas do Sudeste e Centro-Oeste do Brasil

ARTICLES

Floristic relationships among inland swamp forests of Southeastern and Central-Western Brazil¹

Relações florísticas entre florestas paludosas interioranas do Sudeste e Centro-Oeste do Brasil

Aloysio de Pádua Teixeira^*; Marco Antonio Assis

Universidade Estadual Paulista, Instituto de Biociências, Departamento de Botânica, Caixa Postal 199, 13506-900 Rio Claro, SP, Brazil

ABSTRACT

We evaluated the floristic relationships among 20 swamp forests of Southeastern and Central-Western Brazil using multivariate analyses. Detrended correspondence analysis (DCA) and TWINSPAN (Two way indicator species analysis) indicated two distinct floristic groups among forests, according to the Phytogeographic Province (Paranaense or Cerrado) and their climate conditions, phytophysiognomies, and species composition. Within the same province, edaphic conditions and geographical distance, among other factors, may be responsible for similarities or dissimilarities among the forests floras. Our results indicated that, despite the low a diversity, γ diversity is high among the forests, as a result of the low floristic similarities among the remnants and the high number of unique species (55% of all species). Although floristically distinct, we concluded that the inland swamp forests of Southeastern Brazil and the gallery swamp forests of Central Brazil are part of the same forest formation. These forests have in common, in addition to the swampy environment, low plant species diversity and species that have high local densities, such as Calophyllum brasiliense Cambess., Cecropia pachystachya Trécul, Dendropanax cuneatus Decne. & Planch., Guarea macrophylla Vahl, Magnolia ovata (A. St.-Hil.) Spreng., Protium spruceanum (Benth.) Engl. and Tapirira guianensis Aubl.

Key words:Cerrado province (Brazilian savannas), floristic similarity, gallery swamp forests, Paranaense province, γ diversity

RESUMO

As relações florísticas entre 20 florestas paludosas interioranas do Sudeste e Centro-oeste do Brasil foram avaliadas por meio de análises multivariadas. A DCA (Análise de correspondência destendenciada) e a TWINSPAN (Two way indicator species analysis) indicaram dois grupos floristicamente distintos, conforme a província fitogeográfica (Paranaense ou Cerrado) e suas condições de clima, fitofisionomias e composição de espécies. Dentro de uma mesma província, as condições edáficas e a distância geográfica, entre outros fatores, podem ser responsáveis por semelhanças ou dissimilaridades florísticas entre as florestas. Os resultados evidenciam que, apesar da baixa diversidade a, a diversidade γ é alta para essas formações, em função de baixas similaridades florísticas entre os remanescentes e do elevado número de espécies exclusivas (55% do total de espécies). Embora floristicamente distintas, conclui-se que as florestas paludosas interioranas do Sudeste do Brasil e as florestas de galeria inundáveis do Brasil Central são parte de uma mesma formação florestal. Estas florestas apresentam, em comum, além do ambiente paludoso, uma baixa diversidade vegetal e espécies com elevadas densidades locais, como Calophyllum brasiliense Cambess., Cecropia pachystachya Trécul, Dendropanax cuneatus Decne. & Planch., Guarea macrophylla Vahl, Magnolia ovata (A. St.-Hil.) Spreng., Protium spruceanum (Benth.) Engl. e Tapirira guianensis Aubl.

Palavras-chave: diversidade γ, mata brejosa de galeria, Província do Cerrado, Província Paranaense, similaridade florística

Introduction

Swamp forests are unique riverside formations with regard to their floristic composition. Their occurrence is restricted to hydromorphic soils which are permanently saturated throughout most of the year (Leitão Filho 1982). Despite the specificity of the soil in which they occur, these forests are widely distributed in the Neotropics and present an interface with different neighboring forests or grasslands. In Southern and Southeastern Brazil, the swamp forests are linked mainly to different forests of Paranaense and Atlantic phytogeographic provinces, such as the seasonal semideciduous inland forests (Rocha et al. 2005, Teixeira & Assis 2005) and the rainforests of the coastal zone (Scarano 2002, 2006). In the central-northern portion of the state of São Paulo and in most of the state of Minas Gerais, as well as in Central Brazil, the swamp forests occur in swampy parts of gallery forests, which are associated with different phytophysiognomies of the Cerrado (Brazilian savannas), like the cerradão (woody cerrado) and cerrado sensu stricto (Oliveira Filho et al. 1990, Nogueira & Schiavini 2003, Gomes et al. 2004, Guarino & Walter 2005, Teixeira & Assis 2009). Within the same province, such as the coastal zone in Atlantic domain, the swamp forests may occur in different edaphic conditions, such as on sandy, mainly Quaternary alluvium-derived substrates (Scarano 2002) or peat soils (Sztutman & Rodrigues 2002, Dorneles & Waechter 2004).

Swamp forests are restricted to hydromorphic soils that determine their natural fragmentation (Ivanauskas et al. 1997) and consequently expose their borders (Costa et al. 1997). This allows the occurrence of species that are prevalent in the surrounding formations, like the cerrados (Marques et al. 2003, Teixeira & Assis 2009), semideciduous seasonal forests (Toniato et al. 1998, Teixeira & Assis 2005) and rain forests (Lieberman et al. 1985, Scarano 2002) to occur in these forests. The typical species of well-drained soils usually have little influence on vegetation structure as they occur in low densities, but they are responsible for increasing the local richness of the swamp forests (Marques et al. 2003, Scarano 2006, Teixeira et al. 2008) and the floristic dissimilarity among the remnants (Costa et al. 1997, Ivanauskas et al. 1997, Teixeira & Assis 2005).

Although the floristic composition of swamp forests has been the subject of various studies, mainly in the state of São Paulo, very little is known about the floristic relationship among swamp forests in different regions of Brazil. This is a function of three factors: the lack of floristic surveys outside the state of São Paulo, the joint investigation of the flora of these forests together with that of other floristically diverse riverside forests (e.g. Rodrigues & Nave 2000, Silva et al. 2007, Martins et al. 2008) and the floristic evaluation of only a few samples in very restricted geographic portions (e.g. Costa et al. 1997, Teixeira & Assis 2005). For other Brazilian vegetation types, such as seasonal semideciduous forests, rainforests and cerrados, the climatic conditions represented by rainfall, humidity, temperature, existence and duration of a dry season or frost, have been cited as the main factors responsible for floristic and structural variations in regional scales of a few hundred kilometers (Salis et al. 1995, Torres et al. 1997, Oliveira Filho & Fontes 2000, Scudeller et al. 2001, Durigan et al. 2003).

A joint analysis of eight swamp forest remnants in the interior of the state of São Paulo showed that these forests have similar structural characteristics and that it is common to find, in several of the remnants, a concentration of many individuals from a few flood tolerant species. However, the low number of studies used for comparison and the spatial proximity among these samples did not enable the authors to evaluate the existence of phytogeographic patterns mainly related to climatic variables across different regions (Teixeira & Assis 2005), as seen in the other main vegetation types of Brazil. In the present study, a comparison was made of the floristic composition of swamp forests studied in the Southeastern and Central-Western regions of Brazil that present interface with seasonal semideciduous forests or cerrados. Our aim was to investigate the floristic similarities and differences among the swamp forests, considering not only the state of São Paulo, but also other states where studies have been performed. As the surrounding vegetation is considered an important factor that may contribute to the dissimilarity among the remnants, we predicted that the swamp forests that present interface with the cerrados would present greater floristic similarities, compared to the swamp forests present in the area of semideciduous seasonal forests, which in turn would form their own groups.

Material and methods

In order to compare the different swamp forests, we considered 20 studies on the inland of São Paulo (SP) and Minas Gerais (MG) States, as well as those in Central Brazil - Distrito Federal (DF), Mato Grosso (MT) and Mato Grosso do Sul (MS) - covering the Paranaense and Cerrado phytogeographic provinces, according to Cabrera & Willink (1973) (figure 1). In the Cerrado province, the swamp forests are called gallery swamp forests (sensu Ribeiro & Walter 2008). To determine whether a forest was to be included in the analyses, two characteristics had to be met: water saturated soil most of the year and water table near the surface in the dry season.

In the studies, the sampling effort was quite varied, depending on different criteria for inclusion of individuals and the size of the samples (table 1). However, due to the low number of studies found, the sampling effort was not considered a restriction for including a study in the comparisons. For the study in the state of Mato Grosso (Oliveira Filho et al. 1990), held in different parts of a gallery forest, only the swampy portion was considered, described by the authors as "the brejo site lies in a more gently sloping part of the stream where the soils are very swampy". For this portion, we calculated the Shannon diversity index, based on the listing of the density of individuals per species provided in the article.

For the analyses we constructed a presence/absence matrix of the species in each remnant, considering the classification system of the Angiosperm Phylogeny Group (APG II 2003, Souza & Lorenzi 2005). We considered only those taxa identified at species level, which underwent a review of nomenclature to correct synonyms. The matrix was formed by 440 species of shrubs and trees, distributed in 203 genera and 77 families. The floristic similarity among the swamp forests was assessed by the Jaccard coefficient (S_J) and multivariate analyses of ordination and grouping. For the ordination analysis we used the DCA - Detrended Correspondence Analysis (Hill & Gauch 1980) and for the grouping analysis we used TWINSPAN - Two way indicator species analysis (Gauch & Whittaker 1981). The Jaccard coefficient was calculated for each pair of forests by the FITOPAC program (Shepherd 1994), considering the complete floristic list (440 species). For the multivariate analysis, the species occurring in only one remnant (241 or 55% of the total species) were excluded in order to better understand the possible groupings of the samples, according to their most frequent elements. Multivariate analyses were processed by the program PC-ORD version 4.0 (McCune & Mefford 1999).

Results

In the 20 comparative studies, the species most frequently found were Calophyllum brasiliense Cambess., Dendropanax cuneatus Decne. & Planch. and Tapirira guianensis Aubl. (19 studies or 95% of the total), Magnolia ovata (A. St.-Hil.) Spreng. (18 studies or 90%), Cecropia pachystachya Trécul and Protium spruceanum (Benth.) Engl. (17 studies or 85%) and Guarea macrophylla Vahl (14 studies or 70%). The richest genera were Ocotea and Miconia (20 species), Ficus (12), Inga (11) and Myrcia (10), while the families represented by a greater number of species were Fabaceae (48 species), Myrtaceae (39), Lauraceae (35), Melastomataceae (23), Rubiaceae (20), Moraceae (17), Euphorbiaceae (16), Annonaceae (15), Meliaceae (13), and Myrsinaceae (11).

The Jaccard coefficient (S_J) showed that, in general, the floristic similarity among the 20 areas was very low because, of the 380 floristic comparisons, only 15 had values greater than or equal to 0.25 (table 2). The highest similarity values, according to the coefficient, were observed between two forests studied in the Distrito Federal (Riacho Fundo and Acampamento streams, S_J = 0.47), between Jeriquara and Cristais Paulista, in state of São Paulo (S_J = 0.39) and between Acampamento stream and Mata da Onça, both in the Distrito Federal (S_J = 0.39). The forests studied in Mato Grosso, Mato Grosso do Sul and southern Minas Gerais (Coqueiral and Santa Rita de Caldas) showed low similarity coefficients between one another and with all other forests studied, indicating that they had the most unique floras in the samples.

The first division generated by TWINSPAN separated the forests studied mainly in the state of São Paulo from those studied in Triângulo Mineiro (a region in the state of Minas Gerais where Uberlândia is inserted) and Central Brazil (figure 2). As an indicator species, the analysis only pointed to Richeria grandis Vahl for the second group. The most important species in the first group were Ardisia ambigua Mart., Aspidosperma cylindrocarpon Müll. Arg., Calyptranthes concinna DC., Citronella gongonha (Mart.) R.A. Howard, Cecropia glaziovi Snethl., Chionanthus trichotomus (Vell.) P. S. Green, Croton floribundus Spreng., Eugenia florida DC., Ficus obtusiuscula (Miq.) Miq., Geonoma brevispatha Barb. Rodr., Guapira opposita (Vell.) Reitz, Inga marginata Willd., Myrcia laruotteana Cambess., Nectandra nitidula Nees, Ocotea diospyrifolia (Meisn.) Mez, Pera obovata (Klotzch) Baill., Persea venosa Nees & Mart. ex Nees, Podocarpus sellowii Klotzsch ex Endl., Styrax pohlii A. DC., Syagrus romanzoffiana (Cham.) Glassman, Tabebuia umbellata (Sond.) Sandwith, Trichilia claussenii C. DC. and T. pallida Sw., among others. In the second group, the preferential species were Cybianthus glaber A. DC., Emmotum nitens (Benth.) Miers, Erythroxylum amplifolium Baill., Ferdinandusa speciosa Pohl, Gomidesia lindeniana O. Berg, Hirtella glandulosa Spreng., Ilex affinis Gardner, Lamanonia ternata Vell., Maprounea guianensis Aubl., Miconia cuspidata Mart. ex Naudin, Myrcia castrensis (O. Berg) D. Legrand, Protium heptaphyllum (Aubl.) Marchand, Pseudolmedia laevigata Trécul, Sclerolobium paniculatum Vogel, Styrax camporum Pohl, Tabebuia serratifolia (Vahl) G. Nicholson, Tococa formicaria Mart., Virola sebifera Aubl., V. urbaniana Warb., Xylopia aromatica (Lam.) Mart. and X. emarginata Mart., among others. The following more frequent species stood out as non-preferential: Calophyllum brasiliense, Cecropia pachystachya, Guarea kunthiana A. Juss., G. macrophylla, Magnolia ovata, Hyeronima alchorneoides Allemão, Protium spruceanum and Tapirira guianensis, among others.

In the DCA, the eigenvalues for the first two axes were 0.456 (axis 1) and 0.238 (axis 2). These two ordination axes were responsible for explaining 65% of the variance of the data. Axis 1 separated the forests mainly according to the phytogeographic province in which they are inserted (figure 3A). On the left of the diagram prevailed swamp forests that present interface with seasonal semideciduous forests in the states of São Paulo and Minas Gerais. At the other extreme, on the right side, are the swamp forests that occur in the Cerrado province: Uberlândia (state of Minas Gerais), Distrito Federal and state of Mato Grosso. Between the two situations, but closer to the first group, we found the forests studied in the state of São Paulo, located in municipalities with a contact zone between forests and cerrado such as Bauru, Agudos, Itirapina, Cristais Paulista and Jeriquara. The distribution of some species in the remnants can be seen in figure 3 (B-I). The two analyses (TWINSPAN and DCA) were consistent, showing similar groups.

Discussion

The recent increase in the number of studies in swamp forests and the insertion of new locations within the state of São Paulo and outside the state has enabled the evaluation of the floristic differences among the forests. The high number of unique species (55% of a total of 20 studies) suggests that the floristic composition of swamp forests is quite heterogeneous and that each remnant has floristic peculiarities that are defined by a set of factors that vary between and within each phytogeographical province. Confirming the initial expectation, the forests were grouped according to the phytogeographical province in which they are located, indicating the importance of the surrounding matrix (semideciduous forest, cerrado or contact between the semideciduous forest and the cerrado) in the definition of the floristic groupings observed. The main factor that varies between the provinces is the climate, with a predominance of Köppen's Cw in the Paranaense province and Aw in the Cerrado. Among the major climatic differences are the lower average temperatures in the former and increased seasonality in the rainfall of the latter (Oliveira Filho & Ratter 1995).

Although some populations are widely distributed and have high local densities in swamp forests, like Calophyllum brasiliense, Cecropia pachystachya, Guarea macrophylla, Dendropanax cuneatus, Hyeronima alchorneoides, Protium spruceanum and Tapirira guianensis, other species, tolerant to water saturation of the soil, were associated to forests of a given province. For example, Podocarpus sellowii, Styrax pohlii and Tabebuia umbellata were important only in the swamp forests of the Paranaense province and Ferdinandusa speciosa and Richeria grandis were only important in the Cerrado province. Moreover, some species typical of swamp forests of Central Brazil, such as Mauritia flexuosa L. f., Pseudolmedia laevigata and Xylopia emarginata (Oliveira Filho et al. 1990, Guarino & Walter 2005), were also found in the state of São Paulo, but restricted to swamp forests established in the municipalities with a greater or lesser degree of cerrado cover.

Complimentary species, in turn, may also have wide distribution in swamp forests, regardless of the phytogeographical province, or they may be associated with forests from a certain province. Species like Casearia sylvestris Sw., Copaifera langsdorffii Desf., Lacistema hasslerianum Chodat, Myrcia splendens (Sw.) DC., Ocotea corymbosa (Meisn.) Mez, O. pulchella Mart., Pera glabrata (Schott) Poepp. ex Baill., Platypodium elegans Vogel, Rapanea umbellata (Mart.) Mez and Siparuna guianenses Aubl., eventually found in swamp forests, occur in cerrados as well as in seasonal semideciduous and riverside forests (e.g. Costa & Araújo 2001, Nunes et al. 2003, Teixeira et al. 2004, Teixeira & Rodrigues 2006). On the other hand, species like Ardisia ambigua, Aspidosperma cylindrocarpon, Citronella gongonha, Cecropia glaziovi, Croton floribundus, Ficus obtusiuscula, Guapira opposita, Inga marginata, Syagrus romanzoffiana and Trichilia claussenii, preferentially found in the group from swamp forests in the state of São Paulo, are commonly found in forests of the Paranaense province (e.g. Pagano et al. 1987, Oliveira Filho et al. 1994, 2001, Carvalho et al. 1996, Santos & Kinoshita 2003). Species like Emmotum nitens, Hirtella glandulosa, Maprounea guianensis, Sclerolobium paniculatum, Styrax camporum, Virola sebifera and Xylopia aromatica, and other important species from the swamp forests of the Triângulo Mineiro and Central Brazil, are common species in the Cerrado phytophysiognomies, especially the cerradão (e.g. Oliveira Filho & Ratter 1995, 2002). With regard to S. paniculatum, it should be emphasized that this species is currently treated as three different taxa that occur in different habitats in the Cerrado (gallery forests, cerrado sensu lato and transition forests of the Amazon) (Oliveira Filho 2006, Silva & Lima 2007). Although they have low local densities mainly in better drained soils or on the edges of swamp forests, the complementary species can contribute greatly to an increase in the γ diversity (regional) of these forests. Moreover, the preferential species in a given province were probably the most responsible for the low similarity values found among the forests here investigated, as well as the floristic groupings observed.

Within the same province, the spatial proximity, soil and climatic conditions, history of human interventions and stochastic events may be influencing the floristic similarities or differences among the swamp forests. The influence of spatial proximity can be determined by observing the highest rates of similarity found between the forests studied in the same municipality (like Brasília, Distrito Federal) or in adjacent municipalities, such as Jeriquara and Cristais Paulista. However, the floristic similarity between nearby areas is not a rule, as other studies located within the same municipality, such as Campinas or Brotas (state of São Paulo), did not show great floristic similarity amongst one another. Additionally, the influence of altitude can be seen, for example, in the floristic composition of the forest studied in Santa Rita de Caldas (state of Minas Gerais), which is situated about 1,180 m above sea level and is therefore subjected to low temperatures and frost occurrence. This forest contains species typical of Southern Brazil, such as Araucaria angustifolia (Bertol.) Kuntze, Ilex paraguariensis A. St.-Hil. and Dicksonia sellowiana Hook. (Loures et al. 2007).

The forest studied in Bonito (state of Mato Grosso do Sul) had the most distinct soil conditions, since these soils are influenced by carbonate rocks, resulting in the deposition of limestone, high calcium content and alkaline pH. These conditions foster the occurrence of species found in seasonal deciduous forests on calciferous or basalt formations and in the Caatinga province (Constantino 2002). These factors may have influenced the low floristic similarity between this forest and the other forests analyzed.

Considering the floristic comparison among eight swamp forest remnants studied in the interior of São Paulo, Teixeira & Assis (2005) listed 149 species, of which 70% occurred in only one of the remnants. By increasing the analysis by 12 more studies, including those carried out in other states, the number of species increased to 440 in total, 55% of which were restricted to only one remnant. While there may be a tendency towards a decrease in the number of unique species with the increase in the number of studies and while the a diversity is low in most of the remnants, the γ diversity of these forests can be considered high, due to the high number of species, most of them unique, and low floristic similarity among forests. The main factor responsible for increasing the γ diversity is the inclusion of studies from the Cerrado province, with distinct phytophysiognomies, flora and climatic conditions from the Paranaense province. These conditions foster the establishment of different complementary species in swamp forests, supporting the hypothesis that the surrounding vegetation exerts an important contribution to the flora of these forests.

Although distinct floristic composition exist among the swamp forests, with low similarity values and complementary species originating mainly from the predominant matrix in the region, we can conclude that the inland swamp forests of Southeastern Brazil and the swamp forests of Central Brazil, called swamp gallery forests, should be treated as part of the same forest type. Besides the swampy environment, these forests share low plant diversity and species with high local densities, such as Calophyllum brasiliense, Cecropia pachystachya, Dendropanax cuneatus, Guarea macrophylla, Magnolia ovata, Protium spruceanum and Tapirira guianensis. These species should inevitably be used in these formations for forest restoration projects, together with exclusive or preferential species from the phytogeographical province in which the swamp forests are located.

Acknowledgments - We thank Capes for the doctoral scholarship provided to the first author; Fapesp for the research grant (Process 07/52613-5); Bruno MT Walter, Jeanine M. Felfili (in memoriam) and Maria T.Z. Toniato, for sending floristic checklists to add to our database; Leandro J. Fonseca for designing the map; Bruno M.T. Walter and the anonymous referees for their constructive comments on the manuscript.

(received: June 03, 2009; accepted: January 27, 2011)

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1

Part of the thesis of first author, Programa de Pós Graduação em Ciências Biológicas (Biologia Vegetal), Universidade Estadual Paulista , Rio Claro, SP, Brazil.

Publication Dates

• Publication in this collection
  11 Apr 2011
• Date of issue
  Mar 2011

History

• Received
  03 June 2009
• Accepted
  27 Jan 2011